Tuesday, July 14, 2020

[Botany • 2020] Amomum foetidum (Zingiberaceae) • A New Species from Northeast Thailand

Amomum foetidum Boonma & Saensouk

in Boonma, S. Saensouk & P. Saensouk, 2020.
 Taiwania. 65(3).

Amomum foetidum (Zingiberaceae), a new species from Northeast Thailand is here described, illustrated and photographed. The key to three species of Amomum which cited and treated in this paper is provided.

Keyword: Amomum cinnamomeum, Amomum foetidum; New species; Northeast; Thailand; Zingiberaceae


Amomum foetidum Boonma & Saensouk, sp. nov.

Fig. 2. Amomum foetidum Boonma & Saensouk;
 A. dissection (from left): dorsal corolla lobe, lateral corolla lobes, labellum, stigma, stamen, calyx, floral tube with stamen and ovary, flower (side view); B. flower (top and semi-side view); C. flower (top view); D. stigma (front and side view); E. stamen (front, side, and back view); F. ovary with epigynous glands (side and front view).
 Photographed by Thawatphong Boonma.

Fig. 1. Amomum foetidum Boonma & Saensouk;
A. habit; B. ligule-side view; C. ligule-front view; D. leaf-margin with the prickly hairs at 1/3 of the leaf tip; E. leaf-apex; F. rhizome with fibers; G. inflorescence above the ground; H. inflorescences and underground part.
Photographed by Thawatphong Boonma.

Fig. 3. Amomum foetidum Boonma & Saensouk;
A. labellum; B. dorsal corolla lobe; C. lateral corolla lobes; D. calyx; E. stigma (front and side view); F. stamen; G. ovary with epigynous glands (side and front view); H. stamen with corolla tube and ovary; I. habitat with inflorescence and the underground parts; J. close up the inflorescence. Drawn by Thawatphong Boonma.

Distribution: So far known from Sakon Nakhon province and mostly cultivated in Northeastern Thailand. 

Etymology: The specific epithet “foetidum” is derived from the Latin word refers to pungent odor which this plant emits the smell similar to the smell of the Stink Bugs (Tessaratoma papillosa Drury) when sniffing near its flower or crushed the part of this plant. 

  Uses: used to substitute the stink bugs (Tessaratoma papillosa Drury) in cooking. 

Notes: all parts of the plant have a pungent odor similar to the smell of stink bugs (Tessaratoma papillosa Drury), bracts and peduncles are white when young or if it in the underground but it will be green if it above the ground.

Thawatphong Boonma, Surapon Saensouk and Piyaporn Saensouk. 2020. Amomum foetidum (Zingiberaceae), A New Species from Northeast Thailand. Taiwania. 65(3); 364-370. 

[Paleontology • 2020] A Revision of the Early Jurassic Ichthyosaur Hauffiopteryx (Reptilia: Ichthyosauria), and Description of A New Species from southwestern Germany

Hauffiopteryx typicus (von Huene, 1931)

in Maxwell & Cortés, 2020.
 DOI: 10.26879/937  

Hauffiopteryx typicus is an Early Jurassic ichthyosaur species from Europe, for which geographically partitioned morphological variation between specimens from England and Germany has been described. We provide a complete anatomical description of the German material to address this taxonomical issue. We also identify and describe a new species of Hauffiopteryx from the southwest German Basin, Hauffiopteryx altera sp. nov., differring from H. typicus in the morphology of the arrangement of cranial elements surrounding the external nares. A phylogenetic analysis recovers the German and English material referred to H. typicus as sister taxa, suggesting that these are indeed conspecific. H. typicus forms a monophyletic group with H. altera and a specimen from the Pliensbachian of Switzerland previously referred to Leptonectes tenuirostris but consistent with H. typicus. We conclude that Hauffiopteryx represents a valid genus, defined by a set of synapomorphies from both the skull and postcranium. Parsimony analysis recovers Hauffiopteryx as sister taxon to Stenopterygius + Ophthalmosauridae.

Keywords: Hauffiopteryx; new species; Posidonienschiefer Formation; Germany; Toarcian; Early Jurassic

ICHTHYOSAURIA Blainville, 1835
PARVIPELVIA Motani, 1999


Type species. Hauffiopteryx typicus (von Huene, 1931)

Revised generic diagnosis. A small- to mid-sized ichthyosaur, 2-3 m in length, with relatively short and slender antorbital rostrum; slight overbite present (unlike Eurhinosaurus, Excalibosaurus in which a substantial overbite is present); prefrontal participating in margin of external naris (unlike Eurhinosaurus, Leptonectes, Stenopterygius); nasals with a greater contribution to the dorsal midline of the rostrum anterior to the external narial opening than the premaxillae (unlike Stenopterygius); prefrontal with larger external exposure on dorsal skull roof than the postfrontal (unlike Stenopterygius); parietal foramen situated anterior to the supratemporal fenestra (situated at the same level as the anterior edge of the supratemporal fenestra in Stenopterygius); supratemporal fenestra small and circular in dorsal view (similar to Eurhinosaurus but unlike Stenopterygius); supratemporal palmate in posterodorsal view (similar to Eurhinosaurus, unlike Stenopterygius); teeth with smooth enamel (unlike Suevoleviathan); gastralia extending posteriorly to the thirty-fifth presacral vertebra (unique feature among Toarcian ichthyosaurs); rib tuberculum and capitulum widely separated in the dorsal region (unlike Temnodontosaurus, Suevoleviathan, Eurhinosaurus); glenoid end of scapula anteriorly expanded (unlike in Temnodontosaurus); coracoid with anterior notch, foramen between humerus, radius, and ulna absent (unlike Suevoleviathan, Temnodontosaurus trigonodon); anterior digit in both fore- and hindfins with notches on the anterior edge (unlike in Suevoleviathan); proximal limb elements angular and forming a tightly articulated mosaic; ischium and pubis thin and styloidal, fused laterally but widely separated medially (autapomorphic); fibula larger than the tibia; hindfin digits converging distally.

FIGURE 1. Hauffiopteryx typicus. A, lectotype, GPIT 1491/4; B, SMNS 51552; C, SMNS 80226. Part 1B © SMNS / M. Wahler.

FIGURE 2. Hauffiopteryx typicus, cranial morphology.
A–G, overview of cranial morphology. A, lectotype, GPIT 1491/ 4; B, SMNS 51552; C, MHH 9; D, SMNS 80226. E, basioccipital in ventral view (SMNS 80226); F, maxillary and posterior dentary teeth (GPIT 1491/4); G, anterior mandible with displaced teeth (SMNS 80226). Scale bars in cm (parts B, D, E).

 Abbreviations: an, angular; bo, basioccipital; de, dentary; hc, hyoid corpus; hy, hyoid element; en, external narial opening; ex, exoccipital; f, frontal; j, jugal; la, lacrimal; lj, lower jaw; mx, maxilla; n, nasal; op, opisthotic; pa, parietal; pal, palatine; pf, prefrontal; pl, palate; pm, premaxilla; po, postorbital; pof, postfrontal; pt, pterygoid; Q, quadrate; qj, quadratojugal; sa, surangular; scr, sclerotic ring; sp, splenial; sq, squamosal; st, supratemporal; s, stapes; utf, supratemporal fenestra.

FIGURE 4. Postcranial morphology of Hauffiopteryx typicus.
 A, anterior neural spines of the lectotype GPIT 1491/4, note the broad axial neural spine of the same height as the neural spine of C3 but twice as wide. B, Fused atlas-axis complex (SMNS 80226); C, gastralia in the mid-dorsal region of SMNS 80226, the arrow indicates the boomerangshaped medial element (see also inset);
D, reconstruction of Hauffiopteryx typicus; the shape and position of the dorsal and caudal fins are based on the lectotype. Anterior is to the left in all parts. Scale bars in cm (parts B, C).
 Abbreviations: at-ax, atlas-axis complex; ns2, neural spine (axis).

Hauffiopteryx typicus (von Huene, 1931)

Hauffiopteryx altera sp. nov. 
 2005 Eurhinosaurus longirostris (Mantell, 1851): Jäger, p. 29, fig. 23. 

 Etymology. The specific epithet is derived from the Latin altera, which means different from/other and refers to the anatomical divergence from the type species, H. typicus.

We found no evidence that the English and German material referred to the genus Hauffiopteryx represent separate species; however, we do find evidence that two species of Hauffiopteryx were present in the Early Jurassic of the Southwest German Basin: H. typicus and H. altera sp. nov. These two taxa are differentiated primarily based on characters pertaining to the lacrimal, prefrontal, and jugal. In addition, we refer a specimen previously considered to be Leptonectes tenuirostris to Hauffiopteryx typicus, extending the range of the genus into the Pliensbachian. This result is supported by phylogenetic analysis, which recovers Hauffiopteryx as a sister group to Stenopterygius + Ophthalmosauridae. Hauffiopteryx represents a valid genus, defined by a suite of synapomorphies involving both the skull and postcranium. However, the functional and paleoecological significance of these characters differentiating Hauffiopteryx from the superficially similar genus Stenopterygius are unclear.

Erin E. Maxwell and Dirley Cortés. 2020. A Revision of the Early Jurassic Ichthyosaur Hauffiopteryx (Reptilia: Ichthyosauria), and Description of A New Species from southwestern Germany. Palaeontologia Electronica. 23(2); a31. DOI: 10.26879/937 

Plain Language Abstract: Hauffiopteryx is a small ichthyosaur, 2–3 meters in length, with small, pointed teeth and large eyes. It is known from the Early Jurassic (~182 million years ago) of southwestern Germany and England, but is most abundant in the Posidonia Shale of southwestern Germany, a world famous Early Jurassic fossil locality. Hauffiopteryx contains only one named species, H. typicus; however, there has been some debate as to whether the German and English finds are actually different species. We examined the anatomy of the German Hauffiopteryx specimens in detail in order to answer this question. This study revealed a second species of Hauffiopteryx from Dotternhausen, Germany, Hauffiopteryx altera sp. nov., differing from H. typicus in skull characteristics. We found no evidence that the English and German specimens referred to Hauffiopteryx typicus represent different species. Additionally, we identified the oldest record of Hauffiopteryx from the Pliensbachian of Switzerland, more than two million years older than the German and English finds. Hauffiopteryx is closely related to Stenopterygius + Ophthalmosauridae, the lineage of ichthyosaurs that includes some of the last representatives of the group.

Monday, July 13, 2020

[Crustacea • 2020] Nanoplax thomai & Pseudopanopeus pusillus • A New Crab of the Genus Nanoplax from the Gulf of Mexico, and Assignment of Micropanope pusilla to A New Genus (Brachyura, Pseudorhombilidae)

Pseudopanopeus pusillus (A. Milne-Edwards, 1880)

in Felder, 2020. 

Nanoplax thomai n. sp. is described from the Gulf of Mexico, representing the second species of the genus. The description is based upon a number of specimens previously misidentified as Micropanope truncatifrons Rathbun, 1898, including one so represented in recent molecular phylogenetic analyses. As restricted, Micropanope truncatifrons remains known with certainty from only the limited type series, which does not include a mature male, and sequence quality tissues are not available for molecular phylogenetic analyses. Its generic placement remains questionable following morphological study of its type materials and comparisons to specimens representing other present and former members of Micropanope Stimpson, 1871. Those comparisons underscore that morphological and molecular distinctions warrant assignment of Micropanope pusilla A. Milne-Edwards, 1880 to a new genus, herein designated as Pseudopanopeus n. gen.

Keywords: Crustacea, Pseudorhombilidae, Gulf of Mexico, Nanoplax, new species, generic revision

Nanoplax thomai n. sp. 

Etymology. The species is named for Brent P. Thoma in recognition of his substantial contributions to understanding of xanthoid crab phylogenetic relationships and in gratitude for the many ways in which he has facilitated the present author’s work over the last decade

Pseudopanopeus pusillus n. comb.
male, cw 5.6 mm (ULLZ 14362 = USNM 1536353) northeastern Gulf of Mexico;

Pseudopanopeus n. gen. 

Etymology. The prefix “pseudo”, for false, is paired with the suffix “panopeus”, derived from the type genus of Panopeidae Ortmann, 1893. As the gender is masculine, its combination with the type species name becomes Pseudopanopeus pusillus (A. Milne-Edwards, 1880). While resembling panopeids in general habitus and long grouped among members of that family, this genus is one of several such that instead fall among Pseudorhombilidae in the phylogenetic analyses of Thoma et al. (2014). 

Pseudopanopeus pusillus (A. Milne-Edwards, 1880) n. comb.
Micropanope pusilla A. Milne-Edwards, 1880

Darryl L. Felder. 2020. A New Crab of the Genus Nanoplax from the Gulf of Mexico, and Assignment of Micropanope pusilla to A New Genus (Crustacea, Brachyura, Pseudorhombilidae). Zootaxa. 4810(3); 531–545. DOI: 10.11646/zootaxa.4810.3.9

[Paleontology • 2020] Trierarchuncus prairiensis • the Last Alvarezsaurid: Hell Creek Formation (uppermost Maastrichtian), Montana

 Trierarchuncus prairiensis
Fowler, Wilson, Freedman Fowler, Noto, Anduza & Horner, 2020

• A growth series of hand claws and other fossil material from a new taxon, Trierarchuncus prairiensis, of the unusual theropod family Alvarezsauridae is described.
• One claw is the most complete known for Alvarezsauridae and challenges assumptions about the lack of curvature in derived members of the group and demonstrates that alvarezsaurid claws could be of higher curvature than typically reconstructed.
• The possible growth series suggests that alvarezsaurid manual unguals may have changed morphology through ontogeny, with younger individuals retaining more basal characteristics. This has implications for taxonomy and phylogeny of the group as some taxa are described based on probable juvenile remains.
• Trierarchuncus derives from the uppermost Maastrichtian Hell Creek Formation of Montana, and is therefore the youngest known alvarezsaurid and was one of the very last dinosaurs, which went extinct at the K–Pg boundary mass extinction.

The enigmatic Alvarezsauria (Dinosauria: Theropoda) are characterized by extremely short forelimbs with a single functional digit bearing a large, robust ungual. Alvarezsauria are first recorded from the Jurassic of China, but are otherwise mostly known from the Cretaceous of South America and Asia, including a number of relatively complete skeletons. North America has yielded only a fragmentary skeleton from the lower Maastrichtian Horseshoe Canyon Formation, Alberta, and a pubis, partial ischium, and metatarsal from the upper Maastrichtian Hell Creek Formation, Montana, and Lance Formation, Wyoming. Here we describe new alvarezsaurid material from the Hell Creek Formation, Montana, comprising a metatarsal III, distal end of a radius, and three manual digit (MD)-I unguals, which form a progressive size series. One MD-I ungual is the most complete yet described, and is much more curved than typically depicted for Alvarezsauridae. Manual D-I unguals are of particular interest as they undergo a number of changes within the clade, including enclosure of the ventral blood vessel groove, development of a ventral sulcus, and increased robusticity and rugosity. Comparison among the new specimens suggests that these features also develop ontogenetically, which may have taxonomic implications. Stratigraphic data shows that alvarezsaurids occur through most of the ∼85 m thick Hell Creek Formation, with the uppermost specimen occurring ∼10 m below the upper contact with the Fort Union Formation. As such these are the youngest known alvarezsaurid remains and demonstrate that the clade survived at least until ∼1–200 kyrs before the Cretaceous–Paleogene mass extinction.

Keywords: Dinosaur, Theropoda, Alvarezsauridae, Mononykus, Hell Creek Formation, Late Cretaceous

 Systematic Paleontology 
Dinosauria Owen, 1842 
Saurischia Seeley, 1888 

Theropoda Marsh, 1881 
Maniraptora Gauthier, 1986 

Alvarezsauridae Bonaparte, 1991 

Trierarchuncus prairiensis gen. et sp. nov. 

Derivation of name. The genus name refers to the 'trierarch', specifically a seafaring ship's captain (trirēmē ships of ancient Greece), and 'uncus' meaning hook (Latin). Species name means 'of the prairie', referring to the gentle plains of eastern Montana (in particular the American Prairie Reserve) where the new material was discovered.


Denver W. Fowler, John P. Wilson, Elizabeth A. Freedman Fowler, Christopher R. Noto, Daniel Anduza and John R.Horner. 2020. Trierarchuncus prairiensis gen. et sp. nov., the last alvarezsaurid: Hell Creek Formation (uppermost Maastrichtian), Montana. Cretaceous Research. In Press. DOI: 10.1016/j.cretres.2020.104560

Trierarchuncus prairiensis (“Captain Hook of the prairie”): a new species of bizarre hook-handed alvarezsaurid dinosaur.

[Entomology • 2020] Cicindelidia cyanipleura • A New Tiger Beetle (Coleoptera, Cicindelidae) from the Trans-Mexican Volcanic Belt

Cicindelidia cyanipleura 
Duran & Roman, 2020

A new tiger beetle species, Cicindelidia cyanipleura Duran and Roman n. sp., of the tribe Cicindelini, is described from the southwestern section of the Trans-Mexican Volcanic Belt. Superficially, it appears to be most closely related to C. rufiventris (Dejean, 1825) but is distinguished on the basis of multiple morphological characters and behavior. Given the rock-loving natural history of the species, escape behavior, polished pronotum, and small body size, it may be more closely related to the petrophiles C. laetipennis (Horn, 1913) or C. politula (LeConte, 1875), from which it is distinguished on the basis of multiple morphological characters and biogeography.

Keywords: Coleoptera, Cicindelidia, new species, Jalisco, Mexico, taxonomy

FIGURE 4. Cicindelidia cyanipleura n. sp. Frontal habitus.
A) Male. B) Female. Male labrum exhibits a concave margin, medially. Female labrum unidentate, margin not concave medially.

Cicindelidia cyanipleura Duran & Roman, n. sp.
Etymology. Cicindelidia cyanipleura n. sp. is named after the blue sides of the elytra and epipleura, derived from Greek: cyan- = blue, -pleura = side.

Daniel P. Duran and Stephen J. Roman. 2020. A New Tiger Beetle from the Trans-Mexican Volcanic Belt (Coleoptera, Cicindelidae, Cicindelini). Zootaxa. 4810(2); 375–382. DOI: 10.11646/zootaxa.4810.2.11

Sunday, July 12, 2020

[Paleontology • 2020] Lusovenator santosi • A New Carcharodontosaurian Theropod from the Lusitanian Basin: Evidence of Allosauroid Sympatry in the European Late Jurassic

  Lusovenator santosi
Malafaia, Mocho, Escaso & Ortega, 2020

Carcharodontosaurian allosauroids were temporally restricted to the Cretaceous, being known from all land masses with the exception of Antarctica. In addition to Veterupristisaurus from Tanzania, exceptions to this distribution have been reported recently, consisting on fragmentary materials from Upper Jurassic strata of China, Germany, and Portugal. Here, we propose a new Late Jurassic carcharodontosaurian taxon, Lusovenator santosi, gen. et sp. nov. based on the reevaluation of previously described specimens from the Lusitanian Basin, Portugal. The performed phylogenetic analysis recovered Lusovenator santosi as an early branching carcharodontosaurian allosauroid diagnosed by an exclusive combination of characters, including three autapomorphic features: (1) large recesses in neural arch of anterior dorsal vertebrae; (2) well-developed and continuous longitudinal laminae extending from the tip of the prezygapophyses to the distal end of the postzygapophyses in mid-caudal vertebrae; and (3) supraacetabular crest of ilium forming a prominent ventrolaterally projecting shelf. Lusovenator santosi is the oldest carcharodontosaurian allosauroid yet discovered from Laurasia and supports unequivocally the hypothesis of a pre-Cretaceous scenario for the radiation of the clade. The identification of this taxon highlights the high diversity of medium- to large-bodied theropods in the later part of the Late Jurassic of the Iberian Peninsula. Carcharodontosauria is not yet known in correlative levels of the North American Morrison Formation, and the existence of contacts after the late Tithonian between these landmasses could explain the distribution of this clade and other dinosaur groups present in the Iberian Jurassic and in the North American Lower Cretaceous.

FIGURE 1. Lusovenator santosi, gen. et sp. nov., SHN.036, holotype, selected skeletal remains.
A, odontoid and atlantal intercentrum in anterior view. B, cervical vertebra in right lateral view. C, D, anterior dorsal vertebra in C, right lateral and D, ventral views. E, sacral neural arch in posterior view. F, neural arch of anterior caudal vertebra in left lateral view. G, H, middle caudal vertebra in G, dorsal and H, left lateral views. I, right ilium in lateral view; J, K, pubes in J, left lateral and K, ventral views. L, silhouette showing preserved elements of SHN.036. A plus sign (+) indicates a diagnostic character and an asterisk (*) indicates an autapomorphy of Lusovenator santosi, gen. et sp. nov.

Abbreviations: acdl, anterior centrodiapophyseal lamina; act, acetabulum; at in, atlantal intercentrum; bf, brevis fossa; cpf, cupedicus fossa; isp, ischial peduncle; mec, medial crest; ms, medial symphysis; ncs, neurocentral suture; ns, neural spine; od, odontoid; pcdl, posterior centrodiapophyseal lamina; posdf, postzygapophyseal spinodiapophyseal fossa; poz, postzygapophysis; pp, parapophysis; prz, prezygapophysis; pup, pubic peduncle; spof, spinopostzygapophyseal fossa; tp, transverse process. Scale bars equal 25 mm (A), 50 mm (B–H), 100 mm (I–K), and 1 m (L).


THEROPODA Marsh, 1881 
TETANURAE Gauthier, 1986 

CARCHARODONTOSAURIA Benson, Carrano, and Brusatte, 2010 

LUSOVENATOR SANTOSI, gen. et sp. nov.

Etymology— The generic name is composed from the Latin words Luso, referring to Lusitania, the province in Roman Hispania related to the current Portugal, and venator, from the Latin word for hunter; the specific name santosi after José Joaquim dos Santos, who found and collected the holotype.

FIGURE 3. Time-calibrated evolutionary tree for Allosauroidea. Agreement subtree produced in TNT, with additional taxa incorporated (Veterupristisaurus milneri) marked with dashed line (see Fig. S1 for the full version of this tree). The pie charts represent relative frequencies of the clades in different paleogeographic areas calculated based on the number of taxa known in each area. Silhouette of dinosaur modified from the drawing of Scott Hartman. Global paleogeographic reconstructions modified from maps in Blakey (2019).



A new carcharodontosaurian theropod, Lusovenator santosi, gen. et sp. nov., is defined based on a partial skeleton of a juvenile individual, SHN.036, collected in upper Kimmeridgian levels of the Lusitanian Basin. A second specimen, SHN.019, corresponding to some axial and appendicular elements of a large-sized individual, from upper Tithonian strata in the same geographic area, is tentatively proposed as referred material. Also, an isolated femur previously tentatively assigned to Lourinhanosaurus antunesi is interpreted as belonging to a carcharodontosaur. Thus, the fossil record of these allosauroids in the Lusitanian Basin spans from the Kimmeridgian to the upper Tithonian. Lusovenator santosi represents the oldest member of Carcharodontosauria defined in the Upper Jurassic of Laurasia and extends the record of this clade, which was already represented in the Lower Cretaceous of Europe. Despite the great similarity in the theropod faunal composition, the recognition of a carcharodontosaurian allosauroid taxon provides a component of the Upper Jurassic Lusitanian Basin distinct from its contemporaneous Morrison Formation in North America. The development of a terrestrial dispersal route connecting these landmasses after the late Tithonian could explain the absence of Carcharodontosauria in the diverse theropod faunas of the Morrison Formation and, additionally, their presence during the Early and Late Cretaceous of North America. The new carcharodontosaurian theropod provides the first evidence for sympatry among allosauroid theropods in the Late Jurassic of Europe. Additionally, the discovery of L. santosi sheds light on a poorly known period of carcharodontosaurian history and adds important data to the knowledge of the early evolution of these allosauroids.

 Elisabete Malafaia, Pedro Mocho, Fernando Escaso and Francisco Ortega. 2020. A New Carcharodontosaurian Theropod from the Lusitanian Basin: Evidence of Allosauroid Sympatry in the European Late Jurassic. Journal of Vertebrate Paleontology. e1768106.  DOI: 10.1080/02724634.2020.1768106  


[Herpetology • 2020] Limnophis branchi Kissing Cousins: A Review of the African Genus Limnophis Günther, 1865 (Colubridae: Natricinae), with the Description of A New Species from north-eastern Angola

Limnophis branchi
Conradie, Deepak, Keates & Gower, 2020

Photo: Bill Branch.  

The African natricine genus Limnophis is represented by two species: Limnophis bicolor Günther, 1865 and Limnophis bangweolicus (Mertens, 1936). They are stout-bodied, semi-aquatic snakes that mostly feed on fish and amphibians, and occur from Botswana and Namibia in the south throughout most of Zambia and Angola to the Democratic Republic of the Congo in the north. We gathered new material from the ranges of both species in Angola and Zambia in order to examine their taxonomic status and identify any overlooked diversity. We constructed a phylogenetic tree, based on three mitochondrial genes (16S, cytb, ND4) and one nuclear gene (cmos), which includes the first DNA sequence data for Limnophis. Three well-supported lineages were identified, each representing separate species. The taxonomic status of the two currently recognised species is validated, and we describe a new species of Limnophis from north-eastern Angola. The new species is distinguished from the others by the combination of distinct ventral and lateral head colouration and patterning, differences in head and ventral scalation, and uncorrected pairwise genetic distances to both L. bicolor and L. bangweolicus of 5.4–8.1% in cytb, 6.1–8.4% in ND4 and 2.7–8.3% in 16S.

KEYWORDS: Africa, cryptic species, geometric morphometrics, integrative taxonomy, Serpentes, swamp snake

 Limnophis branchi sp. nov. (paratype – PEM R19466)
Photo: Bill Branch. 

Limnophis branchi sp. nov. 
 Branch’s Swamp Snake

Etymology – We name this new snake species in honour of the late William ‘Bill’ Roy Branch (1947–2018), Curator Emeritus of Herpetology at Port Elizabeth Museum, in recognition of his many contributions to African herpetology, especially in Angola during the latter part of his career. The lead author was introduced to reptile systematics and Angolan herpetology by Bill and is grateful for his guidance and mentorship over the years. The name is masculine in gender.

Werner Conradie,V. Deepak, Chad Keates and David J. Gower. 2020. Kissing Cousins: A Review of the African Genus Limnophis Günther, 1865 (Colubridae: Natricinae), with the Description of A New Species from north-eastern Angola. African Journal of Herpetology. DOI: 10.1080/21564574.2020.1782483  
RESUMO: O género africano de natricíneos Limnophis contém duas espécies: Limnophis bicolor Günther, 1865 e Limnophis bangweolicus (Mertens, 1936). Estas cobras semi-aquáticas de corpo robusto alimentam-se principalmente de peixes e anfíbios, e ocorrem no sul de África entre o Botswana e a Namíbia, para norte em grande parte da Zâmbia e Angola até à República Democrática do Congo. Nós colhemos material da área de distribuição de ambas as espécies em Angola e na Zâmbia para examinar o seu estatuto taxonómico, e para identificar se existia alguma diversidade oculta. Construímos uma árvore filogenética baseada em três genes mitocondriais (16S, cytb, ND4) e um gene nuclear (cmos), incluindo os primeiros dados de sequências genéticas para o género Limnophis. Foram identificadas três linhagens bem suportadas, cada uma representando uma espécie distinta. Assim, validámos o estatuto taxonómico das duas espécies já conhecidas, e descrevemos uma nova espécie de Limnophis do nordeste de Angola. Esta nova espécie pode ser distinguida das outras pela combinação de padrões de coloração ventral e lateral da cabeça, e diferenças nas escamas da cabeça e escamas ventrais. As distâncias genéticas não corrigidas entre esta espécie e L. bicolor e L. bangweolicus são de 5.4–8.1% para o gene cytb, 6.1–8.4% para o ND4 e 2.7–8.3% para o 16S.

PALAVRAS-CHAVE: África, morfometria geométrica, taxonomia integrativa, Serpentes, cobra-dos-pântanos

[Mammalogy • 2020] Ancient Divergence Driven by Geographic Isolation and Ecological Adaptation in Forest Dependent Sundaland Tree Squirrels (Genus Sundasciurus)

Sundasciurus spp.

in Hinckley, Hawkins, Achmadi, et al., 2020. 

A surprising amount of hidden phylogenetic diversity exists in the small to medium size, drab colored squirrels of the genus Sundasciurus. This genus is endemic to Sundaland and the Philippines, where it is widespread. An earlier revision of this genus found that the high elevation ‘populations’ of the widespread, lowland slender squirrel (S. tenuis) were different species. Previous phylogenies based on mitochondrial cytochrome b sequences also suggested that the widespread, lowland Low’s squirrel (S. lowii) and the narrow endemic Fraternal squirrel (S. fraterculus) are not reciprocally monophyletic. Additionally, deep divergences have been identified between lineages within Low’s squirrel that date to the early Pliocene. Here we focus on evaluating the relationships and differences within and between populations of these two nominal species using whole mitochondrial genome sequences, nuclear intron sequences, and morphology. We reassess the taxonomy of this group, revalidate the species status of Robinson’s squirrel (Sundasciurus robinsoni Bonhote, 1903) support the species level recognition of the Natuna squirrel (Sundasciurus natunensis Thomas, 1895) and identify three other lineages that require further study. We estimate times of divergence and integrate geologic history to find that most of the divergences are pre-Pleistocene, and thus predate the Pleistocene flooding of Sundaland. Biogeographic, and ecological factors may have played a more important role than climatic factors in generating these patterns. While divergence in allopatry seems to be the main process driving speciation in lowland Sundaland squirrels (Sundasciurus), ecomorphological and behavioral adaptations in this clade suggest an important role of niche divergence.

Keywords: Sundasciurus, mammal, Borneo, biogeography, speciation, systematics, rodent, Sciuridae

Sundasciurus lowii from Sepilok, Sabah.
 photo: Mojito La

Figure 6. Dorsal, palatal, and lateral views of the holotype skulls of Sundasciurus lowii (NHM76.5.2.14), Sundasciurus natunensis (NHM94.9.28.40) and Sundasciurus robinsoni (NHM3.2.6.55). Given the highly damaged state of the holotype skull of Sundasciurus fraterculus (NHM95.1.9.12), the type of Sundasciurus fraterculus siberu (NHM47.1488) is shown instead for the sake of comparison. All pictures are at the same scale.

Figure 1. Map of Sundaland showing the distribution of the different subspecies of the Low-Fraternal squirrel species complex, with dots indicating sample collection locations of specimens included in the morphometric analyses. Samples are colored according to the four highly supported major clades recovered in the mitogenome-based phylogenies of Figures 2, 3. Populations indicated by a red circle have been considered S. l. natunensis but the morphological data in this study suggests they are more similar to S. l. lowii. Populations indicated with a green square have been considered S. l. lowii but the morphological data in this study suggests these they are more similar to S. l. natunensis. Molecular evidence from these populations is needed to clarify their taxonomic status. Populations included in the morphological analyses but not the genetic analyses are indicated in gray.

Sundasciurus robinsoni (Bonhote, 1903) – Robinson’s squirrel

Sciurus robinsoni Bonhote (1903):24.
—Type locality. Bukit Besar, Nawngchik, Patani, Peninsular Thailand, 2500 ft.

Holotype. BMNH, skull, skin, adult female, collected 30 August 1901 by C. H. Robinson and N. Annandale

Subspecific Taxonomy
Sundasciurus robinsoni robinsoni (Bonhote, 1903)
Sciurus robinsoni alacris Thomas, 1908e: 306.—Type locality. Selangor-Pahang border, Malaya, 3000ft.
Sciurus seimundi Thomas and Wroughton, 1909:440.—Type locality. Kundur Island, Riau Islands

Sundasciurus robinsoni balae (Miller, 1903)
Sciurus balae Miller, 1903:14.—Type locality. Tana Bala, Batu Islands, Sumatra
Sciurus piniensis Miller, 1903:14.—Type locality. Pulo Pinie, Batu Islands, Sumatra
Sciurus humilis Miller, 1913:24.—Type locality. Kateman river, East Sumatra

Sundasciurus robinsoni vanakeni (Robinson and Kloss, 1916)
Sciurus vanakeni Robinson and Kloss, 1916:270.—Type locality. Barong Bharu, Korinchi, Sumatra, 4000 ft.

Distribution: Malay Peninsula, Sumatra, and Batu and Rhio Archipelagos. The subspecies S. r. robinsoni and S. r. balae have been recorded from 0–610 m, on the Malay Peninsula and Rhio and Sumatra and Batu, respectively, while S. r. vanakeni has been recorded between 900–1372 m on Mount Kerinci.

Sundasciurus natunensis (Thomas, 1895)- Natuna squirrel.

Sciurus lowii natunensis Thomas, 1895:26.
—Type locality. “Sirhassen Island” (Serasan), Natuna Islands.

Holotype. BMNH, skull, skin, adult male, collected 23 September 1893 by A. H. Everett.

Distribution: Southern Natuna islands (only recorded in Sirhassen Island), and possibly in west Borneo. Despite discrete characters such as postorbital processes and tail shape of the latter populations closely resemble S. natunensis, we currently consider the taxonomic status of west Borneo populations as incertae sedis given the intermediate phenotypic position of these among S. natunensis and S. lowii shown in the PCA and DAPC and the lack of genetic evidence. Genetic data from these, southern Sumatra and northern Natuna populations is needed to clarify the taxonomy and distribution of this group.

Sundasciurus lowii (Thomas, 1895) - Low’s squirrel.

Sciurus lowii Thomas, 1895: 253.
—Type locality. “Lumbidan, on the mainland opposite Labuan”, Sarawak, Borneo.

Holotype. BMNH 76. 5. 2. 14., skull, skin, adult male, collected by H. Low (unknown date).

Subspecific Taxonomy
Sundasciurus lowii lowii (Thomas, 1895)

Sundasciurus lowii bangueyae (Thomas, 1908)
Sciurus lowii bangueyae Thomas, 1908:387.— Type locality. “Banguey Island” (Banggi)

Sundasciurus lowii lingungensis (Miller, 1901)
Sciurus lingungensis Miller, 1901:123.—Type locality. Lingung Island, Near Bunguran Island, Natuna Islands

Distribution: S. l. lowii is present in Borneo, S. l. bangueyae in Banggi and Balambangan islands, (and possibly Malawali island as well) and S. l. lingungensis in the Northern Natunas (recorded in Bunguran, Lingung and Laut islands).

Taxonomic Notes: 
Corbet and Hill (1992) pointed out that geographical variation in Sundasciurus lowii sensu lato is slight. Many of the currently recognized subspecies are not morphologically distinct from others. We consider Sundasciurus robinsoni balae and Sundasciurus robinsoni vanakeni valid subspecies based on their ventral coloration and craniodental differentiation (Figure 5 and Supplementary Figure S3). We synonymize former Sundasciurus lowii humilis with Sundasciurus robinsoni balae due to external resemblance (ventral grayish coloration in limbs). We also synonymize based on external resemblance (lack of ventral grayish coloration in limbs) Sundasciurus lowii seimundi with Sundasciurus robinsoni robinsoni. Finally, we consider Sundasciurus lowii lingungensis and Sundasciurus lowii bangueyae valid subspecies given their craniodental differentiation (Figure 5 and Supplementary Figure S3).

Molecular species delimitation analyses suggest the presence of three species in Borneo within S. lowii (in Sabah, Sarawak and East Kalimantan). However, phenotypic divergence seems to be slight and the fine scale distribution of the morphs is unknown. The only Sarawak sample included in the PCA was clustered separately but close to Sabah + east Kalimantan, that largely overlapped. Regarding the DAPC, only east Kalimantan samples were differentiated from the remaining overlapping populations. Relative size of interorbital breadth seems to differentiate a small number of Sarawak specimens from the remaining S. lowii, but these samples are clustered very close to the other S. lowii. Sarawak specimens also seem to have darker fur than Sabah and East Kalimantan S. lowii and a lack of blond tips on the tail, which are only observed in Sabah populations. We consider these populations unconfirmed candidate species (Padial et al., 2010) until a better molecular and morphological sampling is performed in terms of geographic coverage and number of specimens included.

As pointed out by Hawkins et al. (2016a) the two subgenera, Aletesciurus and Sundasciurus, proposed by Moore (1958) are not supported as reciprocally monophyletic groups. The discrete characters (presence of sagittal crest, skull size, and shape of anterior-mesial lobe of auditory bullae) described as distinguishing subgenera were not valid for all species. For instance, the Palawan mountain squirrel (Sundasciurus rabori) belongs to Aletesciurus but lacks a sagittal crest and has an intermediate antero-mesial lobe and skull size (40.9-43.6 mm) among both subgenera (Heaney, 1979; Hawkins et al., 2016a). The northern Palawan tree squirrel (Sundasciurus juvencus) shows an antero-mesial lobe that resembles that of the subgenus Sundasciurus despite being assigned to Aletesciurus. Finally, different species of the subgenus Sundasciurus such as S. lowii, S. tahan or S. altitudinis have skulls that reach sizes (41–43 mm) that overlap with those of S. rabori (authors unpublished data). Although these discrete characters do not support the subgenera Aletesciurus and Sundasciurus, Heaney (1979) found support for them in his morphometric analyses. Molecular phylogenies suggest that there are 5–6 major phylogenetic lineages within Sundasciurus, which correspond to monophyly in the subgenus Aletesciurus (which contains 3 divergent clades), and paraphyly in the subgenus Sundasciurus (which also contains 2–3 divergent clades) (Den Tex et al., 2010; Hawkins et al., 2016a). The lack of universally valid diagnostic features and the phylogenetic evidence from this and previous studies demonstrate that current Sundasciurus subgeneric classification is invalid, so we synonymize Aletesciurus with Sundasciurus.

Arlo Hinckley, Melissa T. R. Hawkins, Anang S. Achmadi, Jesús E. Maldonado and Jennifer A. Leonard. 2020. Ancient Divergence Driven by Geographic Isolation and Ecological Adaptation in Forest Dependent Sundaland Tree Squirrels. Front. Ecol. Evol. DOI: 10.3389/fevo.2020.00208



[Botany • 2020] Vanilla armoriquensis (Orchidaceae, Vanilloideae) Living in the Clouds: A New High-elevation Species of Vanilla from Peru

Vanilla armoriquensis Damian & Mitidieri 

in Damian & Mitidieri, 2020. 
Illustration by Robb Niklasson

Vanilla armoriquensis, a new species belonging to subgen. Vanilla, is described and illustrated based on living material from Peru. The new species is similar to V. costaricensis but can be easily distinguished by the larger sepals and petals, and a lip with a well-developed triangular apex, bearing a callus with prominent, broad keels, that has an orange coloration at the base and above the middle.

Keywords: Taxonomy; Vanilla costaricensis; V. methonica; V. mexicana; Vanilla subgenus Vanilla; Monocots

Alexander Damian and Nicole Mitidieri. 2020. Living in the Clouds: A New High-elevation Species of Vanilla (Orchidaceae, Vanilloideae) from Perú. Phytotaxa. 451(2); 154–160. DOI: 10.11646/phytotaxa.451.2.5   

[Botany • 2020] Hoya longicalyx (Apocynaceae: Asclepiadoideae) • A New Species of Hoya from Yunnan, China

Hoya longicalyx Wang Hui & E. F. Huang

in Huang, Yang, Xi, et al., 2020. 
長萼球蘭  ||   taiwania.ntu.edu.tw

Hoya longicalyx Wang Hui & E. F. Huang, a new species of Hoya R. Br. (Apocynaceae: Asclepiadoideae) from Yingjiang county (Yunnan, China), is described and fully illustrated. The new species is morphologically similar to H. chinghungensis (Tsiang & P.T. Li) M.G. Gilbert, P.T. Li & W.D. Stevens, but can be easily distinguished by its rounded lamina base, longer and linear calyx lobes, ovate and translucent corona lobes. Result from molecular phylogenetic analysis revealed that the new species was sister to Hoya engleriana Hosseus.

Keyword: Hoya, new taxon, Phylogenic Analysis, ITS, 5’- ETS, psbA-trnH, trnT-trnL, matK

Fig. 3. Hoya longicalyx. A. Habits of the new species in situ, arrows indicate the plants. B. Inflorescence. C. Leaves. D. Branch and inflorescence. E. Leafy stem. F. Petiole. G. Lamina margin. H. Bracteoles. I. Corona (top view and side view). J. Calyx. K. Corolla (adaxial and abaxial sides). L. Pollinarium. M. Pistils. Photographed by H. Wang and E. F. Huang.

Fig. 2. Hoya longicalyx. A. Branch with inflorescences. B. Leafy stem and leaf. C. Bracteole. D. Calyx and ovaries. E. Corolla (adaxial side). F. Corolla (abaxial side). G. Corona (side view). H. Corona lobe (side view). I. Pollinarium. J. Pistils. Drown by Z. M. Li and H. Dong.

Hoya longicalyx Wang Hui & E. F. Huang, sp. nov. 

Type: CHINA. Yunnan Province: Yingjiang county, A-jiang-po, elev. ca. 1900 m, on arboreal branch covered by moss in mid-montane evergreen forest, 16 May 2019, Huang Er-feng 1905003 (holotype SZG!; isotype PE!, KUN!, TAI!). 

Diagnosis: Morphologically similar to Hoya chinghungensis (Tsiang & P.T. Li) M.G. Gilbert, P.T. Li & W.D. Stevens from which it differs in having longer lamina (1.5–2 cm vs. 1–1.5 cm), acuminate lamina apex (vs. acute to obtuse lamina apex), longer calyx lobes (5– 7 mm vs. 1.5–2 mm), ovate and translucent corona lobes (vs. pink and triangular corona lobes) and oblong and upside apart ovaries (vs. ovate and attached ovaries).

Geographical distribution: Hoya longicalyx is known only from the type locality in Yingjiang county, Yunnan province, southwest China, base on the fieldwork and herbaria investigations in HITBC (Herbarium, Xishuangbanna Tropical Botanical Garden, CAS), IBSC (South China Botanical Garden, CAS), PE (Institute of Botany, CAS) and KUN (Herbarium, Kunming Institute of Botany, CAS). 

Ecology: Epilithic on arboreal branch covered by moss (Trachypodopsis serrulata) under mid-montane evergreen forest. 

Etymology: Hoya longicalyx is named from its long and linear calyx lobes, which is a significant feature to distinguish the new species from relevant species.

 Er-Feng Huang, Lei-Lei Yang, Wang Xi, Yi-Ye Liu and Hui Wang. 2020. Hoya longicalyx, A New Species of Hoya (Apocynaceae: Asclepiadoideae) from Yunnan, China. Taiwania. 65(3); 353-359. taiwania.ntu.edu.tw/abstract.php?type=abstract&id=1694